The genus Cladosporium and similar dematiaceous ... - CBS - KNAW

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Crous et al. Fig. 21. Stigmidium schaereri. A. Lichenicolous habit on Dacampia hookeri. B. Vertical section through an ascoma. C–D. Asci. E–G. Ascospores. H. Older, brown ascospores. Scale bars = 10 µm. Stigmidium Trevis., Consp. Verruc.: 17. 1860. (Fig. 21). Type species: Stigmidium schaereri (A. Massal.) Trevis., Consp. Verruc.: 17. 1860. Description: Roux & Triebel (1994, figs 47–50). Notes: The type species of the genus is lichenicolous, characterised by semi-immersed, black, globose ascomata with ostiolar periphyses and periphysoids. Asci are 8-spored, fasciculate, bitunicate, (endotunica not giving a special reaction in Congo red or toluidine blue). Ascospores are fusoid-ellipsoidal, medianly 1- septate, guttulate, thin-walled, lacking a sheath. Presently no culture is available, and thus the placement of Stigmidium remains unresolved. Discussion From the LSU sequence data presented here, it is clear that Mycosphaerella is not monophyletic as previously suggested (Crous et al. 2001, Goodwin et al. 2001). The first step to circumscribe natural genera within this complex was taken by Braun et al. (2003), who separated Cladosporium anamorphs from this complex, and erected Davidiella (Davidiellaceae; Schoch et al. 2006) to accommodate their teleomorphs. The present study reinstates the genus Teratosphaeria for a clade of largely extremotolerant fungi (Selbmann et al. 2005) and foliar pathogens of Myrtaceae and Proteaceae (Crous et al. 2004a, b, 2006b, 2007b), and further separates generic subclades within the Mycosphaerellaceae, while Batzer et al. (2007) again revealed Schizothyrium Desm. (Schizothyriaceae) to cluster within the Mycosphaerellaceae. Our results, however, provide support for recognition of Schizothyrium as a distinct genus, although Schizothyriaceae was less well supported as being separate from Mycosphaerellaceae (Capnodiales). Although pleomorphism represents a rather unstudied phenomenon in this group of fungi, it has been observed in several species. Within the Teratosphaeria clade, Crous et al. (2007b) recently demonstrated teleomorphs to have Readeriella and Cibiessia synanamorphs, while the black yeast genera that belong to this clade, commonly have more than one anamorph state in culture. The present study also revealed Readeriella mirabilis to have two conidial types in culture, and to be highly plastic regarding its mode of conidiogenesis, and Readeriella to be the oldest generic name available for a large group of leaf-spotting coelomycetes in the Teratosphaeriaceae (Capnodiales). Although not commonly documented, there are ample examples of synanamorphs in Capnodiales. Within Mycosphaerella, Beilharz et al. (2004) described Passalora perplexa Beilharz, Pascoe, M.J. Wingf. & Crous as a species with a coelomycete and yeast synanamorph, while Crous & Corlett (1998) described Mycosphaerella stigmina-platani F.A. Wolf to have a Cercostigmina U. Braun and Xenostigmina Crous synanamorph, and recent collections also revealed the presence of a similar species that has typical “Stigmina” (distoseptate conidia) and Pseudocercospora (euseptate conidia) synanamorphs (Crous, unpubl. data), and Crous (1998) reported Readeriella epicoccoides (coelomycete) to 30
Phylogenetic lineages in the Capnodiales have a Cercostigmina (hyphomycete) synanamorph in culture. Although the Mycosphaerella complex encompasses thousands of names, it may appear strange that it is only now that more clarity is obtained regarding the phylogenetic relationships among taxa in this group. This is partly due to the fact that these organisms are cultivated with difficulty, and also that the first paper to address the taxonomy of this complex based on DNA sequence data was only relatively recently published (Stewart et al. 1999). In the latter study, the genus Paracercospora Deighton (scars minutely thickened along the rim), was shown to be synonymous with the older genus Pseudocercospora. Similarily, Crous et al. (2001) showed that Cercostigmina (rough, irregular percurrent proliferations) was also synonymous with Pseudocercospora. This led Crous & Braun (2003) to conclude that conidiomatal type, conidial catenulation, septation and proliferation of conidiogenous cells were of less importance in separating species at the generic level. Mycovellosiella Rangel and Phaeoramularia Munt.-Cvetk. were subsequently reduced to synonymy with the older name, Passalora Fr., and characters identified as significant at the generic level were pigmentation (Cercospora vs. Passalora), scar structure (Passalora vs. Pseudocercospora), and verruculose superficial hyphae (Stenella vs. Passalora). Due to the unavailability of cultures, no decision was made regarding Stenella (verrucose conidia and mycelium), Stigmina (distoseptate conidia), and several other, less well-known genera such as Asperisporium Maubl., Denticularia Deighton, Distocercospora N. Pons & B. Sutton, Prathigada Subram., Ramulispora, Pseudocercosporidium Deighton, Stenellopsis B. Huguenin and Verrucisporota D.E. Shaw & Alcorn. In a recent study, however, Crous et al. (2006a) were able to show that Phaeoisariopsis (synnemata, conidia with slightly thickened hila) and Stigmina (distoseptate conidia) were also synonyms of Pseudocercospora. The present study shows that most anamorph genera are polyphyletic within Teratosphaeria, and paraphyletic within Capnodiales. In some cases, generic concepts of anamorphs based on morphology and conidium ontogeny conform well with phylogenetic relationships, though this is not true in all cases due to convergence. Nevertheless, anamorphs still convey valuable morphological information that is contained in the anamorph name, and naming anamorphs continue to provide a practical system to identify the various asexual taxa encountered. Acknowledgements We gratefully acknowledge several colleagues in different countries who have collected the material studied here, without which this work would not have been possible. We thank M. Vermaas for preparing the photographic plates. M. Grube is gratefully acknowledged for sending us fresh material of Stigmidium to include in this study, and D. Triebel and M. Grube are thanked for advice regarding the strain of hamathecial tissue in Stigmidium. K.A. Seifert is thanked for numerous discussions about anamorph concepts and advice pertaining to this paper, and for recollecting Coccodinium bartschii to enable us to clarify its phylogeny. J.K. Stone is thanked for commenting on an earlier draft version of this paper. References Aptroot A (2006). Mycosphaerella and its anamorphs: 2. Conspectus of Mycosphaerella. CBS Biodiversity Series 5: 1–231. Barnes I, Crous PW, Wingfield BD, Wingfield MJ (2004). Multigene phylogenies reveal that red band needle blight of Pinus is caused by two distinct species of Dothistroma, D. septosporum and D. pini. Studies in Mycology 50: 551–565. Barr ME (1972). Preliminary studies on the Dothideales in temperate North America. Contributions from the University of Michigan Herbarium 9: 523–638. Batzer JC, Mercedes Diaz Arias M, Harrington TC, Gleason ML, Groenewald JZ, Crous PW (2007). Four species of Zygophiala (Schizothyriaceae, Capnodiales) are associated with the sooty blotch and flyspeck complex on apple. Mycologia: in press. Beilharz VC, Pascoe IG, Wingfield MJ, Tjahjono B, Crous PW (2004). Passalora perplexa, an important pleoanamorphic leaf blight pathogen of Acacia crassicarpa in Australia and Indonesia. Studies in Mycology 50: 471–479. Braun U, Crous PW, Dugan F, Groenewald JZ, Hoog GS de (2003). Phylogeny and taxonomy of cladosporium-like hyphomycetes, including Davidiella gen. nov., the teleomorph of Cladosporium s.str. Mycological Progress 2: 3–18. Butin H, Pehl L, Hoog GS de, Wollenzien U (1996). Trimmatostroma abietis sp. nov. (hyphomycetes) and related species. Antonie van Leeuwenhoek 69: 203–209. Castañeda RF, Kendrick B, Gené J (1997). Notes on conidial fungi. XIII. A new species of Cladosporium from Cuba. Mycotaxon 63: 183–187. Chupp C (1954). A monograph of the fungus genus Cercospora. Ithaca, New York. Published by the author. Cortinas M-N, Crous PW, Wingfield BD, Wingfield MJ (2006). Multi-gene phylogenies and phenotypic characters distinguish two species within the Colletogloeopsis zuluensis complex associated with Eucalyptus stem cankers. Studies in Mycology 55: 133–146. Crane JL, Schoknecht JD (1986). Revision of Torula and Hormiscium species. New names for Hormiscium undulatum, Torula equina, and Torula convolvuli. Mycologia 78: 86–91. Crous PW (1998). Mycosphaerella spp. and their anamorphs associated with leaf spot diseases of Eucalyptus. Mycologia Memoir 21: 1–170. Crous PW, Aptroot A, Kang JC, Braun U, Wingfield MJ (2000). The genus Mycosphaerella and its anamorphs. Studies in Mycology 45: 107–121. Crous PW, Braun U (2003). Mycosphaerella and its anamorphs. 1. Names published in Cercospora and Passalora. CBS Biodiversity Series 1: 1–571. Crous PW, Braun U, Schubert K, Groenewald JZ (2007a). Delimiting Cladosporium from morphologically similar genera. Studies in Mycology 58: 33–56. Crous PW, Corlett M (1998). Reassessment of Mycosphaerella spp. and their anamorphs occurring on Platanus. Canadian Journal of Botany 76: 1523– 1532. Crous PW, Denman S, Taylor JE, Swart L, Palm ME (2004a). Cultivation and diseases of Proteaceae: Leucadendron, Leucospermum and Protea. CBS Biodiversity Series 2: 1–228. Crous PW, Groenewald JZ (2006a). Mycosphaerella alistairii. Fungal Planet No. 4. (www.fungalplanet.org). Crous PW, Groenewald JZ (2006b). Mycosphaerella maxii. Fungal Planet No. 6. (www.fungalplanet.org). Crous PW, Groenewald JZ, Gams W (2003). Eyespot of cereals revisited: ITS phylogeny reveals new species relationships. European Journal of Plant Pathology 109: 841–850. Crous PW, Groenewald JZ, Mansilla JP, Hunter GC, Wingfield MJ (2004b). Phylogenetic reassessment of Mycosphaerella spp. and their anamorphs occurring on Eucalyptus. Studies in Mycology 50: 195–214. Crous PW, Kang JC, Braun U (2001). A phylogenetic redefinition of anamorph genera in Mycosphaerella based on ITS rDNA sequence and morphology. Mycologia 93: 1081–1101. Crous PW, Liebenberg MM, Braun U, Groenewald JZ (2006a). Re-evaluating the taxonomic status of Phaeoisariopsis griseola, the causal agent of angular leaf spot of bean. Studies in Mycology 55: 163–173. Crous PW, Palm ME (1999). Systematics of selected foliicolous fungi associated with leaf spots of Proteaceae. Mycological Research 103: 1299–1304. Crous PW, Schroers HJ, Groenewald JZ, Braun U, Schubert K (2006d). Metulocladosporiella gen. nov. for the causal organism of Cladosporium speckle disease of banana. Mycological Research 110: 264–275. Crous PW, Summerell BA, Carnegie AJ, Mohammed C, Himaman W, Groenewald JZ (2007b). Foliicolous Mycosphaerella spp. and their anamorphs on Corymbia and Eucalyptus. Fungal Diversity 26: 143–185. Crous PW, Wingfield MJ, Mansilla JP, Alfenas AC, Groenewald JZ (2006b). Phylogenetic reassessment of Mycosphaerella spp. and their anamorphs occurring on Eucalyptus. II. Studies in Mycology 55: 99–131. Crous PW, Slippers B, Wingfield MJ, Rheeder J, Marasas WFO, Phillips AJL, Alves A, Burgess T, Barber P, Groenewald JZ (2006c). Phylogenetic lineages in the Botryosphaeriaceae. Studies in Mycology 55: 235–253. Doherty MW (1900). New Species of Trimmatostroma. Botanical Gazette 30: 400– 403. Dugan FM, Schubert K, Braun U (2004). Check-list of Cladosporium names. Schlechtendalia 11: 1–103. Ellis MB (1976). More dematiaceous hyphomycetes. Commonwealth Mycological Institute, Kew. Eriksson O (1981). The families of bitunicate ascomycetes. Opera Botanica 60: 1–220. www.studiesinmycology.org 31
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Page 7 and 8: CONTENTS P.W. Crous, U. Braun and J
Page 9 and 10: lectotype for the genus by Clements
Page 11: Schubert K (2005a). Morphotaxonomic
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Page 72 and 73: Arzanlou et al. Table 1. (Continued
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Arzanlou et al. Rhinocladiella mack
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Arzanlou et al. Fig. 26. Veronaea c
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Arzanlou et al. Cultural characteri
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Arzanlou et al. Fig. 30. Myrmecridi
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Arzanlou et al. Fig. 32. Radulidium
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Arzanlou et al. Fig. 34. Rhodoveron
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Arzanlou et al. even further, thoug
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available online at www.studiesinmy
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Dichocladosporium gen. nov. 10 chan
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Dichocladosporium gen. nov. Fig. 3.
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Dichocladosporium gen. nov. to intr
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Dichocladosporium gen. nov. Czechos
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Cladosporium herbarum species compl
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Cladosporium sphaerospermum species
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Crous et al. The aim of the present
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Crous et al. 10 changes Athelia epi
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Crous et al. Table 1. Isolates used
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Crous et al. Table 1. (Continued).
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Crous et al. 0.1 expected changes p
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Crous et al. Fig. 6. Cladophialopho
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Crous et al. Fig. 11. Cladophialoph
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Crous et al. Fig. 13. Cladophialoph
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Crous et al. Fig. 15. Exophiala sp.
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Crous et al. Fig. 18. Cylindrosympo
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Crous et al. Fig. 19. Fusicladium a
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Crous et al. Fig. 22. Fusicladium f
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Crous et al. Fig. 24. Fusicladium p
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Crous et al. Fusicladium rhodense C
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Crous et al. Excluded taxa Polyscyt
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Crous et al. the conidial tips are
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Cladophialophora carrionii complex
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Hormoconis resinae and morphologica
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Fig. 6 Hormoconis resinae and morph
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Cladophialophora, 52 k , 54 k -55,
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Fusicladium phillyreae, 189 c , 191
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Ramichloridium epichloës, 60, 89 P
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Trimmatostroma salicis, 3 t , 5, 6
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